Pressure difference receiving ears.

Directional sound receivers are useful for locating sound sources, and they can also partly compensate for the signal degradations caused by noise and reverberations. Ears may become inherently directional if sound can reach both surfaces of the eardrum. Attempts to understand the physics of such pressure difference receiving ears have been hampered by lack of suitable experimental methods. In this review, we review the methods for collecting reliable data on the binaural directional cues at the eardrums, on how the eardrum vibrations depend on the direction of sound incidence, and on how sound waves behave in the air spaces leading to the interior surfaces of eardrums. A linear mathematical model with well-defined inputs is used for exploring how the directionality varies with the binaural directional cues and the amplitude and phase gain of the sound pathway to the inner surface of the eardrum. The mere existence of sound transmission to the inner surface does not ensure a useful directional hearing, since a proper amplitude and phase relationship must exist between the sounds acting on the two surfaces of the eardrum. The gain of the sound pathway must match the amplitude and phase of the sounds at the outer surfaces of the eardrums, which are determined by diffraction and by the arrival time of the sound, that is by the size and shape of the animal and by the frequency of sound. Many users of hearing aids do not obtain a satisfactory improvement of their ability to localize sound sources. We suggest that some of the mechanisms of directional hearing evolved in Nature may serve as inspiration for technical improvements.

The role of pressure difference reception in the directional hearing of budgerigars (Melopsittacus undulatus).

In many birds, the middle ears are connected through an air-filled interaural pathway. Sound transmission through this pathway may improve directional hearing. However, attempts to demonstrate such a mechanism have produced conflicting results. One reason is that some species of birds develop a lower static air pressure in the middle ears when anaesthetized, which reduces eardrum vibrations. In anaesthetized budgerigars with vented interaural air spaces and presumed normal eardrum vibrations, we find that sound propagating through the interaural pathway considerably improves cues to the directional hearing. The directional cues in the received sound combined with amplitude gain and time delay of sound propagating through the interaural pathway quantitatively account for the observed dependence of eardrum vibration on direction of sound incidence. Interaural sound propagation is responsible for most of the frontal gradient of eardrum vibration (i.e. when a sound source is moved from a small contralateral angle to the same ipsilateral angle). Our study confirms that at low frequencies the interaural sound propagation may cause vibrations of the eardrum to differ much in time, thus providing a possible cue for directional hearing. The acoustically effective size of the head of our birds (diameter 28 mm) is much larger than expected from the dimensions of the skull, so apparently the feathers on the head have a considerable acoustical effect.

Karl von Frisch lecture. Signals and flexibility in the dance communication of honeybees.

Progress in understanding dance communication in honeybees is reviewed. The behaviour of both dancers and follower bees contain flexible and stereotypic elements. The transfer of specific information about direction and distance probably involves more than one sensory modality. The follower bees need to stay behind the dancer (within the angle of wagging) during at least one waggle run in order to perceive the specific information. Within this zone, a small stationary air-flow receiver (like the antenna of a follower bee) experiences a well-defined maximum when the abdomen of the wagging dancer passes by. Within 1 mm from the tip of the abdomen, the maximum may be caused by oscillating flows generated by the wagging motion. At other positions and distances (up to several millimetres from the dancer) the maximum is due to a spatially narrow jet air flow generated by the vibrating wings. The time pattern of these maxima is a function of the angular position of the receiver relative to the axis of the waggle run and thus a potential cue for direction. In addition to the narrow jet air flows, the dancers can generate a broad jet. The jets are not automatic by-products of wing vibration, since they can be switched on and off when the dancer adjusts the position of her wings.

The Tuned Cricket.

Communication by means of sound is not always easy. Sound suffers much attenuation and degradation close to ground. Crickets have adapted to this by exploiting sharply tuned mechanical systems. A coevolution ensures that the calling song of males and the directional hearing of females are tuned to the same frequency.